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2018 Canthidium alvarezi Martínez and Halffter, 1986: a remarkable Ateuchus Weber, 1801 (Coleoptera: : ) Francois Genier Canadian Museum of Nature, [email protected]

Mario Cupello Federal University of Parana, Brazil

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Genier, Francois and Cupello, Mario, "Canthidium alvarezi Martínez and Halffter, 1986: a remarkable Ateuchus Weber, 1801 (Coleoptera: Scarabaeidae: Scarabaeinae)" (2018). Insecta Mundi. 1145. http://digitalcommons.unl.edu/insectamundi/1145

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. July 27 2018 Insecta 0646 1–4 urn:lsid:zoobank.org:pub:D02A2597-5671-4551-A5E7-ACF0C- A Journal of World Systematics 3D330A4 Mundi

0646

Canthidium alvarezi Martínez and Halffter, 1986: a remarkable Ateuchus Weber, 1801 (Coleoptera: Scarabaeidae: Scarabaeinae)

François Génier Research and Collections, Canadian Museum of Nature, PO Box 3443, Station D, Ottawa, Ontario, K1P 6P4, Canada

Mario Cupello Laboratory of Systematics and Bioecology of Coleoptera, Department of Zoology, Federal University of Paraná. Centro Politécnico, Jardim das Américas, 81.531-980, Curitiba, PR, Brazil

Date of issue: July 27, 2018

Center for Systematic Entomology, Inc., Gainesville, FL François Génier and Mario Cupello Canthidium alvarezi Martínez and Halffter, 1986: a remarkable Ateuchus Weber, 1801 (Coleoptera: Scarabaeidae: Scarabaeinae) Insecta Mundi 0646: 1–4 ZooBank Registered: urn:lsid:zoobank.org:pub:D02A2597-5671-4551-A5E7-ACF0C3D330A4

Published in 2018 by Center for Systematic Entomology, Inc. P.O. Box 141874 Gainesville, FL 32614-1874 USA http://centerforsystematicentomology.org/

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Canthidium alvarezi Martínez and Halffter, 1986: a remarkable Ateuchus Weber, 1801 (Coleoptera: Scarabaeidae: Scarabaeinae)

François Génier Research and Collections, Canadian Museum of Nature, PO Box 3443, Station D, Ottawa, Ontario, K1P 6P4, Canada [email protected]

Mario Cupello Laboratory of Systematics and Bioecology of Coleoptera, Department of Zoology, Federal University of Paraná. Centro Politécnico, Jardim das Américas, 81.531-980, Curitiba, PR, Brazil

Abstract. The species Canthidium alvarezi Martínez and Halffter, 1986 (Coleoptera: Scarabaeidae: Scarabaei- nae) is transferred to the genus Ateuchus Weber, 1801, becoming Ateuchus alvarezi (Martínez and Halffter), new combination. Its relationship with other species in the genus is briefly discussed. The validity and ranking of the genus Lobidion Génier, 2010, originally described from a single female, is discussed based on the discovery of male specimens, which lead us to reclassify it as a monotypic subgenus of Ateuchus, becoming Ateuchus (Lo- bidion), new rank, with the only included species becoming Ateuchus (Lobidion) punctatissimus (Génier, 2010) new combination. Key words. Taxonomy, systematics, type specimens, dung , New World. Resumo. Canthidium alvarezi Martínez and Halffter, 1986 (Coleoptera: Scarabaeidae: Scarabaeinae) é trans- ferido para o gênero Ateuchus Weber, 1801, sob a nova combinação Ateuchus alvarezi (Martínez e Halffter). Seu relacionamento com outras espécies no gênero é brevemente discutido. A validade e categorização do gênero Lobidion Génier, 2010, o qual foi descrito a partir de apenas uma única fêmea, é discutida com base em machos recém-descobertos, o que nos leva a considerar o táxon como um subgênero monotípico de Ateuchus, sendo agora tratado como Ateuchus (Lobidion), novo status, com a sua única espécies conhecida se tornando, portanto, Ateu- chus (Lobidion) punctatissimus (Génier, 2010), nova combinação. Palavras-chave. Taxonomia, sistemática, espécimes-tipo, rola-bostas, Novo Mundo.

Introduction The megadiverse New World genera Ateuchus Weber, 1801 and Canthidium Erichson, 1847 (Cole- optera: Scarabaeidae) are currently under investigation respectively by MC and FG. A preliminary assessment of the current species classification for these two genera has shown that several taxa were assigned to the wrong genus, and some should be transferred to different genera (MC, personal observa- tion). The following note results from these observations.

Materials and Methods Specimens studied are deposited in the collection of the Canadian Museum of Nature, Ottawa (CMNC); the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina (MACN) and the Muséum national d’Histoire naturelle, Paris (MNHN). Methodological details concern- ing dissections, genitalic preparation, imaging and plate preparation follow Génier and Moretto (2017). 2 • Insecta Mundi 0646, July 2018 Génier and Cupello

Discussion While organizing the insect collection of the Canadian Museum of Nature, one of us (FG) came across a series of one male and four female specimens labeled as paratypes of “Canthidium superbum Martínez, 1982”. The name is a nomen in litteris and therefore unavailable, but the label data matched the information given for the paratypes of Canthidium (C.) alvarezi Martínez and Halffter, 1986, in the original description. Given that these specimens also match perfectly the description given by Martínez and Halffter (1986) and that the greater part of the Martínez collection is currently housed at the CMNC, these are undoubtedly part of the paratypes studied by Martínez and Halffter (1986). The incongruence between the name appearing on the paratype labels and the actual species name is of little concern as Martínez used to label going-to-be type specimens well before the actual publication of the species descriptions, leading to the existence of several specimens bearing never-published nomina in litteris in the Martínez collection (e.g., “Canthidium violetae”, “Canthon laminatus subandinus” and “Canthon virens nordestinus”). Proper paratype labels, with the name Canthidium alvarezi, are now affixed to the five paratypes currently deposited in the CMNC. The holotype male and the allotype female are deposited in the collection of the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina (Cupello 2018), while two additional paratypes, male and female, are housed in Gonzalo Halffter’s personal collection, Xalapa, Mexico (Martínez and Halffter 1986). As pointed out in the original description, this species is outstanding in appearance by its relatively large size (Fig. 1–3) and by the sinuous pronotal lateral edge in males (Fig. 2A). This modification of the pronotal edge is an adaptation to accommodate the disproportionately elongate femora and tibiae in repose position. In addition to the type series deposited in the Martínez collection (CMNC, MACN), three more individuals, two males and one female, correctly identified by Yves Cambefort, were found in the collection of the Muséum national d’Histoire naturelle in Paris. Two of these specimens are from the Henry Walter Bates collection (MNHN) collected on Mount Roraima. The third specimen has a more precise location being from the Guyanese portion of Mount Roraima (MNHN). This new locality data added to the data from the Martínez collection (viz. Brazil: Roraima [“Rio Branco”]: Pacaraima: Serra de Marari: Surumú; and Venezuela: Bolívar: Canaima National Park), suggest that this species is restricted to the Tepuis region of the Guyana shield (Fig. 6). The overall shape, the green metallic color and the elongate prothoracic legs in the male are features seen in some species of the genus Canthidium. However, upon closer examination, several characters such as the deeply concave anterior hypomeral depression, which is posteriorly bordered by a complete and well-defined carina, and the elytral apex produced into a transverse bulge (Fig. 2B) place this species in the genus Ateuchus Weber as currently defined (Balthasar 1939; Howden and Young 1981; Kohlmann 1997). It should, however, be noted that the most posterior surface of the pronotum possesses a distinct marginal bead on the median third as opposed to the description, which states that the posterior marginal bead is completely lacking. Current observations show that the development of the marginal bead greatly varies in development within the genus as currently defined. Another character, harder to define and never used, the hardness of the integument also places it in Ateuchus. In Ateuchus, the integument is very hard and will sometimes break when a pin is inserted as opposed to the more flexible integument of the species of Canthidium. Consequently, we propose the following nomenclatural change: Ateuchus alvarezi (Martínez and Halffter 1986), new combination. To investigate a possible relationship within the genus Ateuchus, the internal sac of the aedeagus (Fig. 4) was dissected. The naming scheme for the internal sac sclerites follows Tarasov and Génier (2015). Interestingly, the lamella copulatrix, as well as any heavily sclerotized hooks and spicules positioned medially in the internal sac were missing. Only the distal axial+subaxial (A+SA) sclerites complex and an enlarged frontolateral peripheral (FLP) sclerite were present. A very small sclerite was also present adjacent to the FLP, which might be a vestigial superior right peripheral (SRP) sclerite. A few puta- tive closely related species (Ateuchus apicatus (Harold, 1867), A. myrmecophilus (Boucomont, 1935), A. pauperatus (Germar, 1823), A. robustus (Harold, 1868), and A. subquadratus (Harold, 1868)), all of which have elongate prothoracic legs in the male and at least one additional external character similar to those found in A. alvarezi, were dissected for comparison. None of those species presented the reduc- tion of genitalic sclerites observed in A. alvarezi. The only species dissected presenting such reduction is Lobidion punctatissimum Génier, 2010 (Fig. 5). At the time of the description of L. punctatissimum (Génier Canthidium alvarezi: an Ateuchus Insecta Mundi 0646, July 2018 • 3

2010), a single female specimen was known. The morphology of this single individual was so odd that a new genus was erected to accommodate it. Subsequent collecting in 2012–2013 by Patrick Arnaud, Fernando Z. Vaz-de-Mello, and one of us (FG), in the same leaf cutter ant nest on Fazenda São Nicolau (Mato Grosso) yielded 14 more specimens including seven males. Our opinion on the validity and ranking of the taxon has now changed with the discovery of males. It is believed that Lobidion is simply a highly modified Ateuchus as the male head is typical for this genus as presently defined. The compact and more flattened body combined with the greatly enlarged and coarsely striated elytral pseudepipleuron at apex are adaptations to myrmecophily, which are also seen in several other species of Ateuchus and other paracoprid genera such as Ontherus Erichson, 1847 (Génier 1996) and Dendropaemon Perty, 1830 (Génier and Arnaud 2016). For these reasons we lower the rank of Lobidion to a monotypic subgenus under Ateuchus as Ateuchus (Lobidion) new rank and consequently establish Ateuchus (Lobidion) punctatissimus (Génier, 2010) new combination. For differences between Ateuchus s. str. and Ateuchus (Lobidion) the reader is referred to Génier’s (2010) identification key. One of us (MC) is currently work- ing on a comprehensive revision of the genus Ateuchus and it is best to wait for those analyses before formally assigning other species of Ateuchus to the subgenus Lobidion.

Acknowledgments We wish to thank Bert Kohlmann and Andrew B.T. Smith for their critical review of the manuscript. FG wishes to thank Fernando Z. Vaz-de Mello and Patrick Arnaud for companionship while collecting at Fazenda São Nicolau. MC is thankful to his PhD supervisor Prof. Dr. Cibele S. Ribeiro-Costa (Fed- eral University of Paraná, Curitiba, Brazil) for her support and to the Brazilian Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for granting him a PhD scholarship.

Literature Cited Balthasar, V. 1939. Neue Choeridium-Arten (Ins. Col.). 6. Beitrag zur Kenntnis der Scarabaeiden der neotropischen Region. Senckenbergiana 21: 45–66. Cupello, M. 2018. On the types species of the New World dung beetle genus Canthidium Erichson, 1847 (Coleoptera: Scarabaeidae: Scarabaeinae), with an annotated checklist of species. Zootaxa 4388: 451–486. Génier, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Memoirs of the Entomological Society of Canada 170: 1–169. Génier, F. 2010. A review of the Neotropical dung beetle genera Deltorhinum Harold, 1869, and Lobidion gen. nov. (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa 2693: 35–48. Génier, F., and P. Arnaud. 2016. Dendropaemon Perty, 1830: taxonomy, systematics and phylogeny of the morphologically most derived phanaeine genus (Coleoptera: Scarabaeidae, Scarabaeinae, Phanaeini). Zootaxa 4099: 1–125. Génier, F., and P. Moretto. 2017. Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa 4248: 1–110. Howden H. F., and O. P. Young. 1981. Panamanian Scarabaeinae: taxonomy, distribution, and habits (Coleoptera, Scarabaeidae). Contributions American Entomological Institute 18: 1–204. Kohlmann, B. 1997 [1996-1997]. The Costa Rican species of Ateuchus (Coleoptera: Scarabaeidae). Revista de Biología Tropical 44–45: 177–192. Martínez, A., and G. Halffter. 1986. Dos nuevas especies de Canthidium (Coleoptera, Scarabaeidae). Anales de la Escuela Nacional de Ciencias Biologica, México 30: 19–25. Tarasov, S., and F. Génier. 2015. Innovative bayesian and parsimony phylogeny of dung (Coleoptera, Scarabaeidae, Scarabaeinae) enhanced by ontology-based partitioning of morphological characters. PLoS ONE 10(3): e0116671.

Received June 26, 2018; Accepted July 16, 2018. Review editor Paul E. Skelley. 4 • Insecta Mundi 0646, July 2018 Génier and Cupello

Figures 1–6. Ateuchus spp. 1–4) Ateuchus alvarezi. 1) Habitus dorsal, male. 2) Habitus lateral, male. 3) Habitus dorsal, female. 4) Internal sac sclerites. 5) Ateuchus (L.) punctatissimus, internal sac sclerites. 6) Distribution map of A. alvarezi.